Identification of powdery mildew resistance genes in
common wheat (Triticum aestivum L). II. French
cultivars
Fj Zeller, J Lutz, El Reimlein, E Limpert, J Koenig
To cite this version:
Fj Zeller, J Lutz, El Reimlein, E Limpert, J Koenig. Identification of powdery mildew resistance
genes in common wheat (Triticum aestivum L). II. French cultivars. Agronomie, EDP Sciences,
1993, 13 (3), pp.201-207.
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Plant
improvement
Identification of powdery mildew resistance genes
in common wheat (Triticum aestivum L).
II. French cultivars
FJ Zeller
1
J Lutz
1
El Reimlein
E
Limpert
J
Koenig
Technische Universität München, Institut für Pflanzenbau und Pflanzenzüchtung, D-80350 Freising-Weihenstephan,
2
Institut für Pflanzenwissenschaften, Gruppe Phytomedizin, ETH, CH-8092 Zürich, Switzerland;
3
INRA, Station d’amélioration des Plantes, Domaine de Crouelle, F-63039 Clermont-Ferrand, France
(Received
18 October 1991;
accepted
9 November
Germany;
1992)
genes for resistance to the wheat powdery mildew pathogen, Erysiphe graminis f sp tritici, were
in 35 French wheat cultivars and in part in their parents. Cultivars were tested with a set of differential pathogen isolates which had specific interactions for each host line. The results were supplemented by cytological investigations and by consideration of pedigrees. Eighteen cultivars showed susceptible reactions. The remainder formed 9
groups with 1 or 2 resistance genes. Of the 5 known resistance genes present, Pm2 was the most common (in 5 cultivars), followed by Pm4b and Pm6 (each present in 4 cultivars), Pm5 (3 cultivars) and Pm8 (2 cultivars). In addition, an
unknown resistance, tentatively designated Mlar, was identified in 3 cultivars.
Summary — Major
analysed
Triticum aestivum
genes
=
common
wheat /
Erysiphe graminis
f sp tritici
=
wheat
powdery mildew / resistance
Résumé — Identification des gènes de résistance à l’oïdium chez le blé tendre (Triticum aestivum L). II. Cultifrançais. Des gènes majeurs de résistance à l’agent de l’oïdium du blé Erysiphe graminisf sp tritici ont été recherchés chez 35 cultivars français de blé, et quelques-uns de leurs parents. Les cultivars ont été testés avec un ensemble d’isolats différentiels présentant des interactions spécifiques sur chaque lignée hôte. Les résultats ont été
complétés par des observations cytologiques et par une analyse des généalogies. Dix huit cultivars ont montré des
réactions de sensibilité. Les autres forment 9 groupes possédant1 ou 2 gènes de résistance. Des 5 gènes de résistance connus présents chez les cultivars étudiés, Pm2 apparaît le plus fréquemment (chez 5 cultivars) suivi de Pm4b
et Pm6 (chacun présent chez 4 cultivars), Pm5 (3 cultivars) et Pm8 (2 cultivars). De plus, un gène de résistance inconnu a été identifié chez 3 cultivars, nommé provisoirement Mlar.
vars
Triticum aestivum
= blé / Erysiphe graminis f sp tritici = oïdium / gènes de résistance
INTRODUCTION
mildew occurs annually throughout the
wheat growing regions in France and
other
countries. This disease is known to
many
reduce yield significantly. Prior knowledge on the
distribution and efficiency of resistance genes in
current wheat cultivars is necessary, in order to
introduce new genes or to substitute and supplement those which are no longer effective. The
present study deals with the characterization and
identification of mildew resistance in common
wheat cultivars grown in France.
One of the first
common
wheat cultivars in
Europe that showed resistance against powdery
mildew caused by Erysiphe graminis f sp tritici
Powdery
was
common
var
the French cultivar Normandie. This cultihad excellent resistance to individual mildew races in the greenhouse and in field trial
1938; von Rosenstiel, 1938;
Roemer, 1941; Nover, 1941). However, it ap-
(Schlichting,
pears that none of the resistance genes in Normandie was directly transferred into French
wheat cultivars which are currently in agricultural use. Another cultivar used in French breeding programmes was Lontoï from Finland. This
cultivar had a dominant resistance gene as de-
scribed by Goujon (1965). However, it is notable
that with the exception of cultivar Foison the
Finnish cultivar Lontoï is not mentioned in the
pedigrees of currently grown cultivars in France.
Furthermore, several French breeding stations
have been using wheat material of foreign origin
carrying resistance to powdery mildew for decades. In particular, the British wheat cultivar Maris Huntsman which has a very high level of mildew resistance conferred by genes from the
tetraploid wheat species Triticum timopheevii
and the Dutch cultivar Clement, whose mildew
resistance can be traced to the alien variation
derived from rye chromosome 1 R, are prominent
examples. In addition, the French wheat strain
VPM1, first described by Maia (1967) and known
to carry the major resistance gene Pm4b (Doussinault, personal communication) was extensively used in French resistance breeding of common wheat. No systematic inventory of mildew
specific resistance genes has been made in
French wheat cultivars. This paper reports on
the resistance phenotype of 35 recently registered accessions, as well as of their progenitors
and mildew resistance sources.
MATERIAL AND METHODS
The
near-isogenic lines of Chancellor with known
powdery mildew resistance genes (Briggle, 1969) and
line TP 114/2*Starke possessing gene Pm6
(Jørgensen and Jensen, 1972) were kindly provided
by RA Mclntosh, Australia. The remaining differential
cultivars, available French registered cultivars and parent lines were provided by plant breeders and the
genebank of INRA, Clermont-Ferrand, France. Differential mildew isolates of the pathogen used were collected from different parts of Europe (Felsenstein et al,
1991).Mildew tests were performed on primary leaf
segments kept on benzimidazole agar as described by
Lutz et al (1992). Inoculum of the individual isolates
was produced on leaf segments of the highly susceptible German wheat cultivar Nimbus (table I). By using a
settling tower a homogenous spore deposition onto
the plant material was guaranteed (inoculation densities≈ 400-500 spores/cm
). Ten d after inoculation
2
(incubation conditions: 17 °C; continuous light, 10 μE
-2 )
m
-1 disease reactions were assessed on the base
s
of colony number and size. Three main classes of host
reactions were distinguished: r = resistant (0-20% inintermediate (30fection in relation to Nimbus), i
50% infection), s
susceptible (> 50% infection). In
some cases, due to larger variation of disease reactions the creation of combined classes (r, i; i, s) was
necessary. The results obtained are based on at least
3 experiments.
=
=
All cultivars that showed a response pattern indicative of the presence of the resistance gene Pm8 were
cytologically scored with respect to satellited chromosome number in root tip mitoses. As the nucleolus organizer regions of rye are suppressed in the wheat
background it was expected that plants homozygous
for the normal 1 B and 6B wheat chromosomes would
contain 4 major satellited chromosomes, whereas
plants homozygous for the translocated chromosome
pair 1BL/1RS would have only 2 (6B) satellited chromosomes
(Zeller, 1973).
RESULTS
Twelve differential isolates of the pathogen (Lutz
et al, 1992) were used which were able to distinguish between host lines/cultivars with known
genes for resistance (table I). Table II shows the
response patterns of 35 French wheat cultivars.
In addition to a group of susceptible cultivars, 9
groups of mildew resistance could be recognized.
Eighteen cultivars: Beauchamp, Camp-Remy,
Capitole, Champlein, Darius, Festin, Festival, Fidel, Frandoc, Goelent, lena, Milpain, Moisson,
Petrel, Promentin, Recital, Rescler and Scipion
were highly susceptible to most or all of the mildew isolates used in this study, indicating that
of the Pm genes of tableI was present.
Two cultivars, Austerlitz and Berlioz were
found to carry the resistance gene Pm2, as their
reaction corresponded well with the pattern displayed by line Ulka/8*CC (table I). Three cultivars, Gerbier, Pernel and Roazon showed a response pattern indicative of the presence of
gene Pm4b. Another group of 2 cultivars (Talent
and Tarasque, table II) appeared to possess the
same response pattern as Hope and Kormoran
(table I) which are known to carry Pm5 (Heun
and Fischbeck, 1987a,b; Lutz et al, 1992). The
response pattern of Thesée seemed to correspond with gene Pm6 of line TP114/2* Starke
(table I). The pattern of cultivar Voyage, identical
to that of the differential Disponent, is indicative
of the presence of resistance gene Pm8 (table I).
This was confirmed by the presence of only 2 satellited chromosomes instead of 4 in mitotic root
tip cells of Voyage.
none
A group of 3 cultivars, Champtal, Friedland
and Nougat appeared to possess gene Pm2
combined with Pm6. The response pattern of
cultivar Damier was characteristic for the presence of genes Pm2 and Pm8. Cytological analysis of the root tip mitoses in Damier also showed
2 satellited chromosome. Reactions of cultivar
Divio indicated the presence of both genes
Pm4b and Pm5. However, there was a mixed reaction to isolate No 14 with some plants possessing only gene Pm4b.
The cultivars Abo, Aristide and Courtot exhibited reaction patterns characterized by resistance
to isolate No 2 and differing from the patterns of
known major resistance genes.
In addition to the identification of resistance
genes in commercial cultivars the reaction patterns of 7 wheat strains
present in their pedidetermined
(table III). Accessions of
grees
the cultivars Etoile de Choisy, Vilmorin 27, strain
293, Hybrid 40 and US (60) 43 appeared to possess no known major resistance gene. The response pattern of the Finnish wheat cultivar Lontoï which was already described by Goujon
(1965), corresponded well with the pattern of
Ulka/8*CC, indicating the presence of gene Pm2.
Strain VPM1 which was widely used in French
wheat resistance breeding was characterized by
a reaction pattern indicative of the presence of
Pm4b.
were
DISCUSSION
The response patterns of distinct host lines after
inoculation with a collection of differential isolates
of the pathogen have been interpreted on the basis of Flor’s classical gene-for-gene hypothesis
(Flor, 1955). These reactions do not always give
clearcut results. Different background effects and
possible interaction of modifying genes may
complicate interpretation. By using the method
described above it is possible to detect qualitatively inherited gene effects when low disease reactions (<
30% infection) to at least 1 isolate
were obtained. However, quantitative gene effects could not be ruled out in reactions showing
&ap;
infection >
30%. Therefore available information
the response patterns of the corresponding
parent lines, the pedigrees and descendent advanced breeding lines were used in addition to
on
determine the resistance
tested.
gene(s)
in the cultivars
In cultivars Capitole, Champlein, Darius, Festival, Fidel, Frandoc, Moisson, Promentin and
Rescler intermediate reactions to certain isolates
observed. Thus there was no evidence for
known
Pm resistance genes in these cultiany
were
vars.
Gene Pm2 in cultivar Austerliz appears to
have been derived from Argent (table IV), a British wheat strain expressing the same response
pattern (unpublished results). However, it is not
known which parent has donated Pm2 to cultivar
Berlioz.
Resistance gene Pm4b was identified in cultivars Gerbier, Pernel and Roazon (table II). All
these cultivars have strain VPM1 in their pedigree (table IV). VPM1 was first described by
Maia (1967) and is a selection of a cross involving Triticum carthlicum T persicum (table IV). It
is very likely that T carthlicum accessions possess a single gene (Mle, Wolfe, 1963; later designated Pm4b by The et al, 1979) and that this
gene has been introduced into VPM1 (table III;
Doussinault, Mclntosh, personal communica-
Pm5 most probably is cultivar Ducat (table IV),
since Moisson does not appear to possess any
known major resistance gene (table II).
It is remarkable that the resistance in cultivars
Abo, Aristide and Courtot could be detected only
means of isolate No 2. All other isolates
showed susceptible reactions (table II). The resistance is tentatively designated Mlar (Aristide).
Although there is a pedigree relationship between Aristide and Courtot (table IV), the origin
of Mlar is unknown.
by
=
tion).
Cultivars Talent and Tarasque possess gene
Pm5. The origin of this resistance gene could not
be deduced, neither from their pedigrees (table
IV) nor from additional data of some progenitors.
However, cultivars Champlein, Hybrid 40 and Vilmorin 27 can be excluded as possible donors
due to susceptible reactions (tables II, III) as well
as Thatcher (Heun and Friebe, 1990). Moreover,
Talent appears to possess some additional resistance indicated by intermediate reactions to isolates Nos 2 and 15, and, in part, to isolates Nos
5, 6 and 13.
Two cultivars, Voyage and Damier, are characterized by the presence of resistance gene
Pm8, located on the short arm of rye chromosome 1 R. Whether this chromosome segment
arose from a 1 B/1 R wheat-rye translocation or a
whole chromosome (1B)1R substitution requires
further cytological analysis. According to the
pedigree of Voyage, Pm8 must come from line
RPB 72-85a (table IV) since Armada (table I)
carries only gene Pm4b. It appears that Damier
inherited Pm2 from Maris Huntsman and Pm8
from Clement, possessing the 1B/1R translocation or substitution.
It is evident that cultivars Champtal, Friedland
and Nougat possess the gene combination Pm2
and Pm6. This combination can be traced to
Maris Hunstman (table I) which is in the pedigree
of all 3 cultivars (table IV). However, Thesée
which also originated from a cross with Maris
Huntsman carries only Pm6 as revealed by intermediate reactions to corresponding isolates.
Cultivar Divio, which carries 2 resistance
genes, Pm4b and Pm5, may have inherited
Pm4b from strain VPM1 (table III). The source of
Up to the present,
at least 6
major genes have
breeding for mildew
been used in French wheat
resistance. These are mainly genes also used in
resistance breeding in Czechoslovakia (Lutz et
al, 1992) in Germany (Heun and Fischbeck,
1987a,b; Schneider et al, 1991), in Scandinavian
countries (Hovmøller, 1989) and in Switzerland
(Winzeler et al, 1991).
There is little genetic diversity of wheat mildew
resistance in Europe. In addition, the genes used
far provide little protection against the contem-
so
pathogen populations (Limpert et al,
1987; Felsenstein et al, 1991). Therefore novel
sources of resistance are urgently needed. They
porary
be found in common wheat itself, in cultivated and wild Triticum relatives, such as emmer
wheat, Einkorn, Aegilops and rye. Moreover, it is
necessary to consider improved strategies to
make better use of the valuable resources.
can
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